{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T01:51:29Z","timestamp":1791510689759,"version":"4.3.4"},"reference-count":43,"publisher":"Elsevier BV","license":[{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/tdm\/userlicense\/1.0\/"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.elsevier.com\/legal\/tdmrep-license"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-017"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-012"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,11,1]],"date-time":"2026-11-01T00:00:00Z","timestamp":1793491200000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-004"}],"funder":[{"DOI":"10.13039\/100009659","name":"Guangzhou Medical University","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100009659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["82403028"],"award-info":[{"award-number":["82403028"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["elsevier.com","sciencedirect.com"],"crossmark-restriction":true},"short-container-title":["Life Sciences"],"published-print":{"date-parts":[[2026,11]]},"DOI":"10.1016\/j.lfs.2026.124711","type":"journal-article","created":{"date-parts":[[2026,9,25]],"date-time":"2026-09-25T23:44:14Z","timestamp":1790379854000},"page":"124711","update-policy":"https:\/\/doi.org\/10.1016\/elsevier_cm_policy","source":"Crossref","is-referenced-by-count":0,"special_numbering":"C","title":["UFSP2 suppresses UFMylation of NONO to govern glycolysis in hepatocellular carcinoma"],"prefix":"10.1016","volume":"405","author":[{"given":"Xiaohong","family":"Xia","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yujie","family":"Xiang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenshuang","family":"Sun","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liping","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zixuan","family":"Wang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuanfei","family":"Deng","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoqing","family":"Guan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hongbiao","family":"Huang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuning","family":"Liao","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiangyu","family":"Zhang","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"78","reference":[{"issue":"3","key":"10.1016\/j.lfs.2026.124711_bb0005","first-page":"209","article-title":"Global cancer statistics 2020: GLOBOCAN estimates of incidence and mortality worldwide for 36 cancers in 185 countries","volume":"71","author":"Sung","year":"2021","journal-title":"CA Cancer J. Clin."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0010","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.cld.2022.08.004","article-title":"Hepatocellular carcinoma: new developments","volume":"27","author":"Ganesan","year":"2023","journal-title":"Clin. Liver Dis."},{"issue":"10127","key":"10.1016\/j.lfs.2026.124711_bb0015","doi-asserted-by":"crossref","first-page":"1301","DOI":"10.1016\/S0140-6736(18)30010-2","article-title":"Carcinoma Bruix hepatocellular","volume":"391","author":"Forner","year":"2018","journal-title":"Lancet"},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0020","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1038\/s41416-022-01771-0","article-title":"Impact of overweight and obesity on the risk of hepatocellular carcinoma: a prospective cohort study in 14.3 million Koreans","volume":"127","author":"Jun","year":"2022","journal-title":"Br. J. Cancer"},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0025","doi-asserted-by":"crossref","first-page":"126","DOI":"10.1186\/s13046-020-01629-4","article-title":"Emerging roles and the regulation of aerobic glycolysis in hepatocellular carcinoma","volume":"39","author":"Feng","year":"2020","journal-title":"J. Exp. Clin. Cancer Res."},{"issue":"9","key":"10.1016\/j.lfs.2026.124711_bb0030","doi-asserted-by":"crossref","DOI":"10.1016\/j.autrev.2023.103393","article-title":"Posttranslational modifications in psoriatic arthritis: a systematic literature review","volume":"22","author":"Calixto","year":"2023","journal-title":"Autoimmun. Rev."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0035","doi-asserted-by":"crossref","first-page":"90","DOI":"10.1038\/s41392-020-0196-9","article-title":"Regulating tumor suppressor genes: post-translational modifications","volume":"5","author":"Chen","year":"2020","journal-title":"Signal Transduct. Target. Ther."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0040","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s13045-020-01027-5","article-title":"Regulation of PD-L1 expression in the tumor microenvironment","volume":"14","author":"Yi","year":"2021","journal-title":"J. Hematol. Oncol."},{"issue":"12","key":"10.1016\/j.lfs.2026.124711_bb0045","doi-asserted-by":"crossref","first-page":"974","DOI":"10.1016\/j.tcb.2019.09.005","article-title":"The UFMylation system in proteostasis and beyond","volume":"29","author":"Gerakis","year":"2019","journal-title":"Trends Cell Biol."},{"issue":"21","key":"10.1016\/j.lfs.2026.124711_bb0050","doi-asserted-by":"crossref","first-page":"5040","DOI":"10.1111\/febs.16730","article-title":"A guide to UFMylation, an emerging posttranslational modification","volume":"290","author":"Millrine","year":"2023","journal-title":"FEBS J."},{"issue":"8","key":"10.1016\/j.lfs.2026.124711_bb0055","doi-asserted-by":"crossref","first-page":"5256","DOI":"10.1074\/jbc.M610590200","article-title":"Two novel ubiquitin-fold modifier 1 (Ufm1)-specific proteases, UfSP1 and UfSP2","volume":"282","author":"Kang","year":"2007","journal-title":"J. Biol. Chem."},{"issue":"17","key":"10.1016\/j.lfs.2026.124711_bb0060","doi-asserted-by":"crossref","first-page":"4085","DOI":"10.1093\/nar\/21.17.4085","article-title":"Purification and cDNA cloning of HeLa cell p54nrb, a nuclear protein with two RNA recognition motifs and extensive homology to human splicing factor PSF and Drosophila NONA\/BJ6","volume":"21","author":"Dong","year":"1993","journal-title":"Nucleic Acids Res."},{"issue":"2","key":"10.1016\/j.lfs.2026.124711_bb0065","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1016\/S0014-5793(02)03447-6","article-title":"PSF and p54nrb\/NonO\u2013multi-functional nuclear proteins","volume":"531","author":"Shav-Tal","year":"2002","journal-title":"FEBS Lett."},{"issue":"9","key":"10.1016\/j.lfs.2026.124711_bb0070","doi-asserted-by":"crossref","first-page":"3989","DOI":"10.1093\/nar\/gkw271","article-title":"The DBHS proteins SFPQ, NONO and PSPC1: a multipurpose molecular scaffold","volume":"44","author":"Knott","year":"2016","journal-title":"Nucleic Acids Res."},{"issue":"4","key":"10.1016\/j.lfs.2026.124711_bb0075","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1002\/wrna.1280","article-title":"PSF: nuclear busy-body or nuclear facilitator?","volume":"6","author":"Yarosh","year":"2015","journal-title":"Wiley Interdiscip. Rev.: RNA"},{"key":"10.1016\/j.lfs.2026.124711_bb0080","first-page":"1","article-title":"WBP11 inhibits UFL1-mediated UFMylation of NONO to drive hepatocellular carcinoma progression","author":"Huang","year":"2025","journal-title":"Oncogene"},{"issue":"51","key":"10.1016\/j.lfs.2026.124711_bb0085","doi-asserted-by":"crossref","first-page":"20314","DOI":"10.1073\/pnas.0707999105","article-title":"A coactivator trap identifies NONO (p54nrb) as a component of the cAMP-signaling pathway","volume":"104","author":"Amelio","year":"2007","journal-title":"PNAS"},{"issue":"13","key":"10.1016\/j.lfs.2026.124711_bb0090","doi-asserted-by":"crossref","first-page":"4863","DOI":"10.1128\/MCB.02144-06","article-title":"Transcriptional activity of androgen receptor is modulated by two RNA splicing factors, PSF and p54nrb","volume":"27","author":"Dong","year":"2007","journal-title":"Mol. Cell. Biol."},{"issue":"11","key":"10.1016\/j.lfs.2026.124711_bb0095","doi-asserted-by":"crossref","first-page":"2189","DOI":"10.1016\/j.biocel.2009.04.011","article-title":"Transcriptional activity of the murine retinol-binding protein gene is regulated by a multiprotein complex containing HMGA1, p54nrb\/NonO, protein-associated splicing factor (PSF) and steroidogenic factor 1 (SF1)\/liver receptor homologue 1 (LRH-1)","volume":"41","author":"Bianconcini","year":"2009","journal-title":"Int. J. Biochem. Cell Biol."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0100","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1261\/rna.2213506","article-title":"p54nrb is a component of the snRNP-free U1A (SF-a) complex that promotes pre-mRNA cleavage during polyadenylation","volume":"12","author":"Liang","year":"2006","journal-title":"Rna"},{"issue":"7","key":"10.1016\/j.lfs.2026.124711_bb0105","doi-asserted-by":"crossref","first-page":"1197","DOI":"10.1210\/me.2010-0517","article-title":"Consensus PP1 binding motifs regulate transcriptional corepression and alternative RNA splicing activities of the steroid receptor coregulators, p54nrb and PSF","volume":"25","author":"Liu","year":"2011","journal-title":"Mol. Endocrinol."},{"issue":"14","key":"10.1016\/j.lfs.2026.124711_bb0110","doi-asserted-by":"crossref","first-page":"1779","DOI":"10.1101\/gad.1565207","article-title":"The multifunctional protein p54nrb\/PSF recruits the exonuclease XRN2 to facilitate pre-mRNA 3\u2032 processing and transcription termination","volume":"21","author":"Kaneko","year":"2007","journal-title":"Genes Dev."},{"issue":"4","key":"10.1016\/j.lfs.2026.124711_bb0115","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1016\/S0092-8674(01)00466-4","article-title":"The fate of dsRNA in the nucleus: a p54nrb-containing complex mediates the nuclear retention of promiscuously A-to-I edited RNAs","volume":"106","author":"Zhang","year":"2001","journal-title":"Cell"},{"issue":"5","key":"10.1016\/j.lfs.2026.124711_bb0120","doi-asserted-by":"crossref","first-page":"1592","DOI":"10.1073\/pnas.1213317110","article-title":"NONO couples the circadian clock to the cell cycle","volume":"110","author":"Kowalska","year":"2013","journal-title":"PNAS"},{"issue":"5722","key":"10.1016\/j.lfs.2026.124711_bb0125","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1126\/science.1107373","article-title":"PERIOD1-associated proteins modulate the negative limb of the mammalian circadian oscillator","volume":"308","author":"Brown","year":"2005","journal-title":"Science"},{"issue":"18","key":"10.1016\/j.lfs.2026.124711_bb0130","doi-asserted-by":"crossref","first-page":"3027","DOI":"10.1007\/s00018-012-0973-x","article-title":"Paraspeckle nuclear bodies\u2014useful uselessness?","volume":"69","author":"Nakagawa","year":"2012","journal-title":"Cell. Mol. Life Sci."},{"issue":"24","key":"10.1016\/j.lfs.2026.124711_bb0135","doi-asserted-by":"crossref","first-page":"4167","DOI":"10.1038\/s41388-021-01848-9","article-title":"Nuclear scaffold protein p54nrb\/NONO facilitates the hypoxia-enhanced progression of hepatocellular carcinoma","volume":"40","author":"Shen","year":"2021","journal-title":"Oncogene"},{"issue":"23","key":"10.1016\/j.lfs.2026.124711_bb0140","doi-asserted-by":"crossref","DOI":"10.1002\/advs.202301983","article-title":"The conserved LncRNA DIO3OS restricts hepatocellular carcinoma stemness by interfering with Nono-mediated nuclear export of ZEB1 mRNA","volume":"10","author":"Hou","year":"2023","journal-title":"Adv. Sci."},{"issue":"14","key":"10.1016\/j.lfs.2026.124711_bb0145","doi-asserted-by":"crossref","first-page":"1896","DOI":"10.1038\/s41388-017-0069-z","article-title":"Growth arrest and apoptosis induction in androgen receptor-positive human breast cancer cells by inhibition of USP14-mediated androgen receptor deubiquitination","volume":"37","author":"Liao","year":"2018","journal-title":"Oncogene"},{"issue":"27","key":"10.1016\/j.lfs.2026.124711_bb0150","doi-asserted-by":"crossref","DOI":"10.1002\/advs.202302025","article-title":"HSP90\u03b2 impedes STUB1-induced ubiquitination of YTHDF2 to drive sorafenib resistance in hepatocellular carcinoma","volume":"10","author":"Liao","year":"2023","journal-title":"Adv. Sci."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0155","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1038\/s41421-019-0092-z","article-title":"USP10 modulates the SKP2\/Bcr-Abl axis via stabilizing SKP2 in chronic myeloid leukemia","volume":"5","author":"Liao","year":"2019","journal-title":"Cell Discov."},{"issue":"2","key":"10.1016\/j.lfs.2026.124711_bb0160","doi-asserted-by":"crossref","first-page":"92","DOI":"10.1038\/s41388-024-03209-8","article-title":"The Ku70-SIX1-GPT2 axis regulates alpha-ketoglutarate metabolism to drive progression of prostate cancer","volume":"44","author":"Huang","year":"2025","journal-title":"Oncogene"},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0165","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1186\/s11658-024-00665-8","article-title":"SENP3 mediates deSUMOylation of SIX1 to promote prostate cancer proliferation and migration","volume":"29","author":"Shao","year":"2024","journal-title":"Cell. Mol. Biol. Lett."},{"issue":"12","key":"10.1016\/j.lfs.2026.124711_bb0170","doi-asserted-by":"crossref","first-page":"5926","DOI":"10.7150\/thno.56490","article-title":"ZEB1 enhances Warburg effect to facilitate tumorigenesis and metastasis of HCC by transcriptionally activating PFKM","volume":"11","author":"Zhou","year":"2021","journal-title":"Theranostics"},{"issue":"12","key":"10.1016\/j.lfs.2026.124711_bb0175","doi-asserted-by":"crossref","first-page":"1315","DOI":"10.1158\/2159-8290.CD-16-0615","article-title":"Targeting cancer metabolism: dietary and pharmacologic interventions","volume":"6","author":"Vernieri","year":"2016","journal-title":"Cancer Discov."},{"issue":"1","key":"10.1016\/j.lfs.2026.124711_bb0180","doi-asserted-by":"crossref","DOI":"10.1186\/s13046-025-03343-5","article-title":"NONO interacts with nuclear PKM2 and directs histone H3 phosphorylation to promote triple-negative breast cancer metastasis","volume":"44","author":"Li","year":"2025","journal-title":"J. Exp. Clin. Cancer Res."},{"issue":"2","key":"10.1016\/j.lfs.2026.124711_bb0185","doi-asserted-by":"crossref","first-page":"762","DOI":"10.1093\/nar\/gky1166","article-title":"RNF8 mediates NONO degradation following UV-induced DNA damage to properly terminate ATR-CHK1 checkpoint signaling","volume":"47","author":"Deshar","year":"2019","journal-title":"Nucleic Acids Res."},{"issue":"23","key":"10.1016\/j.lfs.2026.124711_bb0190","doi-asserted-by":"crossref","DOI":"10.1002\/advs.202301983","article-title":"The conserved LncRNA DIO3OS restricts hepatocellular carcinoma Stemness by interfering with NONO-mediated nuclear export of ZEB1 mRNA","volume":"10","author":"Hou","year":"2023","journal-title":"Adv. Sci."},{"issue":"5","key":"10.1016\/j.lfs.2026.124711_bb0195","first-page":"1602","article-title":"Metabolic reprogramming and its clinical implication for liver cancer","volume":"78","author":"Yang","year":"2023","journal-title":"Hepatology"},{"issue":"3","key":"10.1016\/j.lfs.2026.124711_bb0200","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1038\/s41568-020-00320-2","article-title":"The metabolism of cancer cells during metastasis","volume":"21","author":"Bergers","year":"2021","journal-title":"Nat. Rev. Cancer"},{"issue":"21","key":"10.1016\/j.lfs.2026.124711_bb0205","article-title":"Erratum: dysregulation of PD-L1 by UFMylation imparts tumor immune evasion and identified as a potential therapeutic target (Proceedings of the National Academy of Sciences of the United States of America (2023) 120 e2215732120)","volume":"120","author":"Zhou","year":"2023","journal-title":"Proc. Natl. Acad. Sci. USA"},{"issue":"9","key":"10.1016\/j.lfs.2026.124711_bb0210","doi-asserted-by":"crossref","first-page":"1056","DOI":"10.1038\/s41556-020-0559-z","article-title":"UFMylation maintains tumour suppressor p53 stability by antagonizing its ubiquitination","volume":"22","author":"Liu","year":"2020","journal-title":"Nat. Cell Biol."},{"key":"10.1016\/j.lfs.2026.124711_bb0215","doi-asserted-by":"crossref","DOI":"10.1016\/j.canlet.2026.218770","article-title":"The GCN5L2-SIX1 complex drives glycolysis to promote progression of triple-negative breast cancer","volume":"659","author":"Liao","year":"2026","journal-title":"Cancer Lett."}],"container-title":["Life Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0024320526005205?httpAccept=text\/xml","content-type":"text\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/api.elsevier.com\/content\/article\/PII:S0024320526005205?httpAccept=text\/plain","content-type":"text\/plain","content-version":"vor","intended-application":"text-mining"}],"deposited":{"date-parts":[[2026,10,9]],"date-time":"2026-10-09T01:27:04Z","timestamp":1791509224000},"score":1,"resource":{"primary":{"URL":"https:\/\/linkinghub.elsevier.com\/retrieve\/pii\/S0024320526005205"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,11]]},"references-count":43,"alternative-id":["S0024320526005205"],"URL":"https:\/\/doi.org\/10.1016\/j.lfs.2026.124711","relation":{},"ISSN":["0024-3205"],"issn-type":[{"value":"0024-3205","type":"print"}],"subject":[],"published":{"date-parts":[[2026,11]]},"assertion":[{"value":"Elsevier","name":"publisher","label":"This article is maintained by"},{"value":"UFSP2 suppresses UFMylation of NONO to govern glycolysis in hepatocellular carcinoma","name":"articletitle","label":"Article Title"},{"value":"Life Sciences","name":"journaltitle","label":"Journal Title"},{"value":"https:\/\/doi.org\/10.1016\/j.lfs.2026.124711","name":"articlelink","label":"CrossRef DOI link to publisher maintained version"},{"value":"article","name":"content_type","label":"Content Type"},{"value":"\u00a9 2026 Elsevier Inc. All rights are reserved, including those for text and data mining, AI training, and similar technologies.","name":"copyright","label":"Copyright"}],"article-number":"124711"}}